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A living Tesseropora (Cirripedia; Balanomorpha) 
from Bermuda and the Azores; first records 
from the Atlantic since the Oligocene 



William A. Newman and Arnold Ross 



ABSTRACT.— Tesseropom occupies a central position in the evolution of the Tetraclitinae, 
and likely the Tetraclitellinae, by virtue of geological age and comparative morphology. 
Considering the biogeography of the two extant species from the Indo-West Pacific, the genus 
is apparently favored by the special req\iirements related to perpetuation of insular populations. 
The more highly evolved genus Tetraclita, on the other hand, is for the most part prevalent on 
continentfd shores where the opportunities for competitive interactions are greater. Recognition 
of a new species of Tesseropora from Bermuda and the Azores corroborates the primarily 
insular nature of the genus and enhances the concept that oceanic iskmds can act as refugia 
for ancient forms. 

In consideration of the special adaptations needed by marine organisms to 
perpetuate their populations on oceanic islands on one hand, and the isolation, 
ephemerality, short faunal lists and concomitant unbalanced biotas of oceanic 
islands on the other, one would expect vagile, eurytopic t£ixa, or the descendents 
thereof, to be disproportionately prevalent on them. Many such t2ixa are early 
members of lineages having advanced members more conmionly found in centers 
of distribution frequently contiguous with continental shores, and this is appar- 
ently the basis for the concept that insular situations can act as refugia for 

ancient forms. 

Tesseropora isseli (de Alessandri, 1895: 296), from the Oligocene of Italy, is the 
oldest known tetraclitid. Pilsbry (1916: 259), when reflecting on the significance of 
the single row of parietal tubes in the waU of the only extant species of the genus 
known at the time, remarked " . . . [T. rosea (Krauss, 1848)] is to be regarded as 
an unprogressive form, which retains characters of the ancestral stock of the 
genus, elsewhere found [in the Tetrachtidae] only in an early stage of develop- 
ment", a fact noted earlier by Darwin (1854:336). Since Pilsbry, Tesseropora has 
maintained a more or less central position in considerations of the evolution of the 
tetraclitine and tetraclitelline branches of the family (Zullo, 1968:274; Ross, 
1969:239; Newman and Ross, 1976:47). 

From a biogeographic point of view, Tesseropora as previously known would 
be considered primarily an insular Indo-Pacific genus; T. rosea occurring solely 
in the southern hemisphere from the southern tip of South Africa to southern 
Australia, the Kermadec Islands and New Caledonia, and T. wireni (Nilsson- 
Cantell, 1921: 366) for the most part in the northern hemisphere, from 
Dar-es-Salaam east, from Chagos Bank to Wake and perhaps the Hawaiian 
Islands. The counterpart of Tesseropora on continental shores is the more 
advanced genus Tetraclita, Recognition of an Atlantic Tesseropora, on Bermuda 
and the Azores, is therefore of considerable significance not only in extending 
the modem distribution of the genus, but also in corroborating biogeographical 
inferences that oceanic islands can act as refugia for ancient forms (see Fig. 4). 



SAN DIEGO SOC. NAT. HIST., TRANS. 18(12):207-216, 24 FEBRUARY 1977 



208 



perfamily Coronuloidea Leach 
Family TetracUtidae Gruvel, 1 
ubfamily Tetraclitinae Gruvel, 

Tesseropora Pilsbry, 1916 

TesseroDora atlantica n. sl 



Tetraclita porosa: Verrill, 1901:22 (Bermuda). 



talactift 



Henry, 



(Bermuda); Ross 



Werner, 1967:70; 



Ross 



(Azores 



Diagno 



A small (diameter of largest spec, approx. 10mm), white 



Tesseropora with moderately developed radii having eroded and (or) oblique 
summits concomitant with a somewhat flaring, toothed orifice; scutal adductoi 
ridge in line and nearly continuous with articular ridge; intermediate articles oi 
cirrus VI supporting five pairs of setae; crest of labrum lacking obvious teeth 



De script 



white or with a slight pinkish cast; conical; radii and 



moderately well developed with summits parallel to base; commonly, when 
eroded, orifice moderately toothed (Fig. 1). Surface of solid radii marked by trans- 
verse ridges, and of paries by low longitudinal ribs equal to the number of longi- 
tudinal tubes in the wall. Sheath white, adpressed, continuous with interior 
surface of paries; region below sheath smooth or marked by reminiscences of small 
basal denticles in the form of numerous fine ribs. Longitudinal septa normal to 
inner and outer laminae, forming a single row of irregular tubes (Fig. 2d); one or 
more incomplete septa may be present on inner surface of outer lamina. None of 
the specimens available is sufficiently eroded to expose the secondary fiUing of 
the parietal tubes, but grinding revealed a white filling near the apex. No calcare- 
ous spines extend from the inner surface of the outer lamina into the parietal 
tubes as reported for wireni by Henry (1957: 33), and as we have observed in 
rosea, but older and larger specimens might have them. Basis calcareous, solid, 
thick, adhering strongly to the wall. Opercular valves white. Scutum with well 
defined adductor muscle pit and ridge, the curvature of the latter being virtually 
continuous with that of the articular ridge (Fig. 2b); rostral and lateral depressor 
muscle crests well defined in larger specimens, apparently increasing in number 
with age; rostral depressors essentially lacking in small specimens. Tergum with 
material forming beak undifferentiated from rest of valve and readily broken or 
worn away; spur furrow open, broad and shallow. 

Trophi undistinguished except for the lack of conspicuous teeth on the labrum 
(Fig. 3g). Cirral counts for two specimens from Bermuda follow: 



I 



II 



III 



IV 



V 



VI 



12 
6 



8 
9 



7 



10 



14 
16 



16 
18 



17 
18 



Specimen 1 



12 

8 



8 

8 



8 
9 



14 
14 



16 
18 



16 
17 



n 

6 



6 

7 



7 



7 



12 
16 



16 
17 



16 




Specimen 2 



12 
6 



6 
7 



7 
9 



12 
14 



15 
17 



18 
19 



1 



superfamilial 



Newman and Ross 



209 





•» 




> 




<^ 



Figure 1. Tesseropora atlantica n. sp., on an oyster from Argus Tower, Bermuda. 

The third cirrus is normal in form (rami not antenniform); supporting three types 
of bipectinate setae (Fig. 3c-e), the heaviest (card) being rarely observed; surfaces 
of the lesser curvature eirmed with short, curved spines or denticles (Fig. 3f). 
Intermediate articles of cirrus VI bearing five pairs of setae with occasional pairs 
or small clusters of short bristles between major pairs (Fig. 3b). The intromittant 
organ is normal. 






B 






B 




Figure 2. A, Right tergum and scutum of Tesseropora wireni (Nilsson-Cantell) from Wake Island 
(type locality of T. w. pacificoy approx. 20mm basal diam.); and B, of T. atlantica n. sp. fr< 
Bermuda (type, approx* 11mm diam.); C, D, and E. basal margins of parietes of T. wireni, T. atlantica 



•111 



and very young Tetraclita stalactifera (Lamarck), respectively. 



210 



B 












1 . 2 111 

--J 1 .Inn 

iCDEFA' 

iHI I.OSm 



I.OSm 



Figure 3. A^, intermediate articles of sixth cirrus in Tesseropora wireni (approx. 20mni diam.) and 
A , portion of an intermediate article of the sixth cirrus in T. wireni (approx. 40mm diam.) from Wake 
I., the latter showing the increase in number of small spines below each of the larger two pairs of 
setae. B-I Tesseropora atlantica n. sp.: B, intermediate articles of sixth cirrus; C, D, and E, specialized 
setae; F, spines found on the third cirrus; G, labrum (right palp deleted); H, first maxilla, I, mandible. 

Type material .—Gxxmet Rock, Bermuda (type locality), intertidal with 



Chthamalus and Catophragmus (see Henry, 1958); USNM 
type); BM(NH) 1976.1281 (paratype) [USNM 
ace. no. 



827 (type lot; USNM 
Argus Tower, southwest of Bermuda, subtidal (2-3m), dried. 



NM 



Lopha frons), W 
168473. UrzeUna 




1976, BM(NH) 



Azores Expedition (see Baker 

Reference 



;e, Azores, 
Southward 



BM(NH) 



makin 



material .—One often must rely on published descriptions in 
_ ments on the distinctiveness of a species. By good fortune 
Tesseropora is a small genus, and we had comparative materials in our collections 
of the other species upon which to base our analysis. 

Tesseropora wireni (Nilsson-Cantell): 

Wake Island [type locality of T. w. pacifica (PUsbry)], numei^ous large speci- 
mens with Euraphia intertexta (Darwin), 1854, R. McFarlan coll., 27 Dec. 1964. 

Ulupau, Hawaii, numerous small specimens from 6" pipe of seawater system. 



Burch leg., 28 Sept 



see discussion 



Moen Island, Truk, CaroUne Islands, a single small speci 
men from the underside of an intertidal coral boulder resting on sand, W. A. New- 



man 



Dublon Channel, Truk, CaroUne Islands, numerous small specimens oi 
from underside of buov, W. A. Newman coll., June, 1956 (see Newman 



pi. 20, fig. 1). 



Table 1. Distinguishing characteristics of extant species of Tesseropora. 



211 







Color: 

1-wall 












Intermediate 


Species 


Form and size 


2-sheath 


Radu 


Parietal 








aegmente 


m 




Sfiiling 




tubes 


Scutum 


Labrum 


Cirrus III 


Cirrus VI 






4-valves 
















Steeply corneal 


1- white 






adductor 


apparently 


occasionally 


five pairs 




when young; 


2- white 






ridge in 


without 


armed with 


of setae 


atlantica 


? when old; 


3-white 


moderately 


in single 


line with 


conspicuous 


heavy bipecti- 


per article 




approx. 10 mm 


4 -white 


wide 


row 


articular 


teeth 


nate setae 






in basal 








ridge 




(cards) 






diameter 








adductor 


with several 


armed with 


three pairs of 






1-white 






ridge 


teeth on 


numerous 


setae per 


m 




2-dirty white 




ditto 


overlapping 


each side of 


cards 


article, num- 


rosea 


Steeply coni- 


tinted pink 


ditto 




articular 


shallow 




erous short 




cal; up to 


3-usu£dly 






ridge 


notch 




setae in dense 




30mm 


pink 
4-white 

1-white 










armed with 


bunches below 
two major prs. 

as rosea l>ut 




Steeply conical 


2-usuaUy 




into sec- 






bipinnate but 


with a few 




when young; 


dark red- 


extremely 


ondary and 


ditto 


ditto 


not bipectinate 


short setae in 


wireni 


low when old; 


brown 


narrow to 


tertiary 






setae (cards) 


groups below 




up to 40mm 


3-usuaUy 
pink 


lacking 


rows bas- 
aUy 








two major 
pairs 






4-white 















Kwajalein Atoll, Marshall Islands, Carmarsel Expedition (CRS-325), inter- 
tidal, on iron piling, numerous specimens with E. intertexta, W. A. Newman coll., 

2 April 1967. 

Ulul Island, Namonuito Atoll, Caroline Islands, Carmarsel Expedition 
(CRS-307), several specimens in Heliopora, with Lithoglyptes wilsoni Tomlinson, 
1969, 10m, W. A. Newman coll., 15 Feb. 1967. 

Port of Palau, Arakabesan Passage near Perir}^! Island, Caroline Islands, in 
Heliopora, Seto Marine Biological Laboratory no. 81, F. Hiro coll., 1934 (see Hiro, 
1935: 5; the collection studied consists of some fragments of Tesseropora, and 
several complete specimens of a pyrgomatid imbedded in the cored). 

Tesseropora rosea (Krauss): 

Gerroa (south of Port Kembla), New South Wales, Australia, on mid-tidal 

rocks, E. Pope coll., 2 April 1963. 

Bouvail-Baie des Tortues, New Caledonia, a single large specimen, dried, 

from a rock, J. C. Plaziat coll., 5 Nov. 1973. 



DISCUSSION 

Only one species referrable to Tesseropora (Conia rosea Krauss) was known 
when Darwin (1854) published his incomparable monograph on the sessile 
barnacles. The second, T. isseli (de Alessandri, 1895), from the Oligocene of Italy, 
while morphologically close to rosea, is known only by the wall and need not con- 
cern us here. A third species, wireni, was described by Nilsson-Cantell (1921: 366), 
followed by wireni pacifica (Pilsbry, 1928: 312) and wireni africana (Nilsson- 
Cantell, 1932: 14). Henry (1957: 33) elevated pacifica to specific rank. We believe, 
as Zullo (1968) seemingly did, that there is presently insufficient data to dis- 
tinguish pacifica from wireni at the specific or the subspecific level, and believe 
the same is true of africana. Therefore, we tentatively treat living Tesseropora as 
containing but two taxa, rosea and wireni. 

The new species, T. atlantica, can be distinguished from both rosea and 
wireni by a number of characters (Table 1), the most obvious being the alignment 
of the scutal adductor ridge with the articular ridge rather than passing well in- 
ward of it (compare Figs. 2b and 2a), and the intermediate segments of cirrus VI 
bearing five rather than three pairs of setae (compare Figs. 3b and 3a). Tesseropora 
rosea, known from New South Wales, South Africa and New Caledonia, differs 
from mature wireni in retaining a single row of parietal tubes throughout life. 
While secondary riblets develop on the interior of the outer lamina, as noted by 



212 



40* — 



E 180" W 




140' 



Tesseroplax 



60' 



40 



O 



20' 



0' 



-20' 



40' 



140' 



20 



E 180' W 



140' 



Figure 4. Distribution of Tesseropora spp. and the closely related Tesseroplax unisemita (Zullo, 1968). 
The latter is known only from Pliocene sediments in the Gulf of CaUfomia (cf. Ross, 1969). The other 
significant fossil record is that for Tesseropora isseli from the Oligocene of Italy, The Indo-West 
Pacific species, T rosea and T wireni, while morphologically distinct, are more closely related to each 
other than to T. atlantica from Bermuda and the Azores. At the present level of our knowledge the 
taxonomic status of the so-called subspecies of T. wireni cannot be sustained. The assignment of 
the Hawaiian form to T wireni is questioned herein. 



Darwin (1854: 336), they are normal to the surface and thereby fail to join the 
primary septa to form secondary tubes. Young wireni, and it is young specimens 
that are apparently most commonly collected and described, also have a single 
row of essentially square parietal tubes, but with growth the longitudinal septa 
join some of the riblets on the interior of the outer lamina, singly or in pairs, 
leading: to the dendritic pattern of the septa separating the secondary tubes 



formed in 



Although we have not observed secondary 



in atlantica, Henry (1958: 224) noted a few in the rostrum of the largest specimen 



IIU 



re like 



she examined. The appearance of large, eroded atlantica may appear 
that of wireni than that of rosea. 

Young, uneroded rosea were described by Darwin (1854: 323), and apparently 
the best character that might be used to distinguish them from young wireni 
would be moderate to well developed radii in the former. It follows that young 
uneroded rosea cannot be distinguished from atlantica, at the present state of our 
knowledge, since both have radii. The only characters 



%%l% 



iwn that will 
separate them are the interiors of the scuta, the pinkish rather than white filling 
of the parietal tubes, and the nature of the cirri described above. 

The specimens of Tesseropora from Hawaii are not tygicsl of wireni. All 
available are small, less than 3 mm in basal diameter, and therefore they had not 



contained larvae and 



comoleted their develooment 



ually mature (see discussion of larvae below). This 



Hawaiian material is similar to the wireni studied 



•]|| 



in having much 



reduced radii and in having a toothed labrum, but the adductor ridge of the 
scutum is closer to being in line with the articular ridge (similar to atlantica), the 
sheath is white (Pilsbry, 1928: 313, noted that the sheath of wireni ranges from 
Prussian blue to white, but his material included specimens from Necker Island in 
the Hawaiian Archipelago as well as from Wake Island), the cirral coimts are 
lower, cirrus III has bipectinate (cards) as well as bipinnate setae, and the inter- 
mediate segments of cirrus VI bear three, four or five pairs of setae (on the same 
ramus), but commonly four, a niunber intermediate between that found in atlan- 
tica. gmd wireni and rosea. The differences noted here make it difficult to assign 



this material to wireni for, if consistent, they would be sufficient to allow c 
propose the Hawaiian form as a new species. However, such a decision will 
to await further studv. 



213 



Both rosea and wireni can attain basal diameters approaching 40 mm, and 
in most large specimens the outer lamina of the shell is all but completely eroded, 
exposing the generally pinkish, more resistent material lining the interior of the 
parieted tubes. In a specimen of rosea from New Caledonia most of these run con- 
tinuously from the apex to the base, but a few can be observed on the exterior, 
beginning more than halfway down the wall rather than at the apex, and this indi- 
cates, although there is predominantly a single row of tubes in the wall, a few 
supernumerary ones occasionally develop. Zullo (1968: 272) noted comparable 
development in this species from southeast Austredia. In appe£U"8ince, wireni is 
comparable, but many more of the pillars exposed by erosion, other than the 
primary ones, begin at varying distances down the shell below the apex. Thus, 
the general ribbing appears finer because of the more numerous development of 
secondeiry parietal tubes. None of the specimens of atlantica is sufficiently eroded 

to expose the pillars. 

It is perhaps significant that wireni, although ranging from East Africa to 
Wake Island and perhaps to Hawaii, appears to be a relatively rare species. Also, 
most of the material described has been of relatively young uneroded specimens, 
up to but generedly smedler than 15 mm or so in diameter. One is led to suspect 
the same thing may be true for atlantica — the specimens on hand are relatively 
young individuals and the species is likely not confined solely to Bermuda and 
the Azores. 

« — 

Verrill (1901: 22) identified a Bermudan form as Tetraclita porosa 
(= squamosa). Although we cannot prove he had Tesseropora rather than 
Tetraclita s. s., it does seem highly likely because he stated, "This is the common, 
small, sessile barnacle foimd on the rocks between tides, with the general appear- 
ance of some species of Balanus". Species of Tetraclita, however, are generally 
relatively large, and it is the erosion of the outer lamina of the shell, exposing the 
colored infilling of the parietal tubes, that gives them the characteristic tetracUtid 
appearance. On the other hand, atlantica is known as a small species, and in being 
uneroded it does have a somewhat balanid appearance. In any event, his identifi- 
cation apparently led Henry (1958: 224) to consider the Stephensons' material to 
be T squamosa stalactifera, the only "subspecies" of the squamosa complex 
known from the Caribbean. The single row of tubes in the wall did not invaUdate 
this conclusion for, as Darwin (1854: 323) noted, in very young specimens of 
Tetraclita there is only a single row of tubes (Fig. 2e). During growth, these are 
added to by bifurcation of the septa at the outer lamina very early in Tetraclita 
and Tetraclitella, quite a little later in wireni and possibly atlantica, and hardly at 
all in rosea. It is important to note that in Tetraclita the septa forming the single 
row of tubes are at angles other than normal to the inner lamina, but in Tesser- 
opora they are normal or essentially normal (compare Figs. 2c and 2d with 2e). 
This and the presence of a calcareous basis in the latter serves to distinguish the 

two genera. 

The development of radii, a tubiferous wall and a strong calcareous basis 
would appear to be fundamental to the Tetraclitidae, all three being the principal 
advances in the shell made over the bathylasmatid (ancestral) level of organization 
(Newman and Ross, 1976: 20). Radii greatly strengthen the waU, important in the 
surf zone (Darwin, 1854: 56; Barnes, Read and Topinka, 1970: 82). A tubiferous 
wall, especially when secondarily filled, places an additional barrier to erosion and 
boring organisms (Ross, 1970: 9; Newman and Ross, 1971: 159). A calcareous 
basis conveys an advantage in forming a strong attachment (Newman, Zullo and 
Wainwright, 1967: 170) and in retarding desiccation in intertidal forms, especially 
in the tropics where porous reef limestones are common ( Southward and Newman, 
in press). Tesseropora has these advanced features. But the most successful 



214 



tetraclitid today, in terms of being an abundant intertidal dominant, is Tetraclita. 
The genus has carried the filled tubiferous wall to the extreme, and perhaps the 
great thickness achieved has allowed it to give up radii, but why it has also given 
up a calcareous basis is an enigma, unless it has gained a degree of limited 
motility, as was demonstrated for Semibalanus balanoides by Crisp (1960: 1208). 

primarily an insular species, is known to carry 



tracli 



imbryonic development through to the cyprid stage, passing the naupliar 



Cantell 



Ross, 1961: 211). Cyprid larvae are 
are not known to remain long in the 



incapable of feeding and, because they also 
plankton, they are not likely propagules for long-range disperal (Newman and 
Tomlinson, 1974: 208). The direct production of the non-feeding stage in diyisa 
could be an adaptation to the relatively sterile waters of most oceanic situations, 
but because most balanomorphans of such regions apparently have nauplii, this 
could be only part of the reason, another being the selection for mechanisms that 
favor maintaining populations on isolated oceanic islands too distant to be regu- 
larly reached from elsewhere by ordinary means. Species having a tendency to 
suppress the nauplius would thereby be likely candidates for maintaining popu- 
lations on isolated insular situations. It is interesting therefore to note that the 
larvae of Tesseropora from Hawaii just mentioned are released as cyprids, and it 
will be interesting to see if the life histories of other populations or species of the 



are so modified 



may aid in maintaining 



insular populations, such populations would appear to have removed themselves 
from the list of potential island colonizers by larval propagules. Furthermore, 
because islands are ephemeral in terms of geological time relative to continental 
shores, the time to extinction of such lineages would be substantially shortened. 
However, this problem can and apparently has been circumvented because some 
sedentary species with short range larvae have achieved and apparently maintain 
distributions over great expanses of open water. So far as divisa is concerned, 
and the likewise virtually cosmopoUtan Balanus trigonus, transport of adult 



gul 



more so by other organisms, seems likely. One of us 



(W.A.N.) has found adults of the former washed ashore on Majuro Atoll m the 
Marshall Islands attached to the shell of the pelagic barnacle Lepas, and of the 
latter, attached to the shell of the whale barnacle Coronula. These species there- 
fore have the potential of being transported great distances as adults, and this 
may explain their virtually cosmopolitan distributions in the warm seas of 



the world. 



known to form an obligate commensal relationship 



individuals of some species are foimd on other organisms, the most notable being 
wireni, which occurs embedded in the blue coral Heliopora. If the history of the 
genus continues and it is eventually completely replaced as a free-living form, 
conceivablv wireni could Dersist as an obligate commensal. 



ACKNOWLEDGMENTS 

We thank Alan J. Southward, Marine Biological Association of the United Kingdom, for 
bringing this problem to our attention and for providing specimens from Bermuda and the Azores. 
We would also like to thank T. E. Bowman, National Museum of Natural History, William K. 
Emerson, American Museum of Natxiral History, W. D. Hartman, Yale Peabody Musuem and W. E. 
Sterrer, Bermuda Biological Station for their attempts to locate voucher specimens from Bermuda 
collected either by Verrill or others near the turn of the century; and G. A. Boxshall, British 
Museum (Natural History) for attempts to locate material from the Azores. Thanks are also due 
Huzio Utinomi, who kindly arranged for the loan of specimens from the Seto Marine Biological 
Laboratory, Beatrice L. Bxirch for the somewhat enigmatic specimens from Hawaii, and Gayle 



Kidder for technical assistance and preparation of the figures. Work supported , in part, by the National 



Science Foundation (DEB-7517149). 



REFERENCES 



215 



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inmiensalism between 




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, F., 1937. Cirripeds of the Palao Islands. 
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